Intensity Of Sexual Selection Research Articles

Female's courtship threshold allows intruding males to mate with reduced effort.

Female decision rules can influence the nature and intensity of sexual selection on males, but empirical demonstrations of rules underlying choice are rare. We hypothesized that female choice is largely based on a courtship duration threshold in the Australian redback spider (Latrodectus hasselti) because females kill males before copulation is complete (premature cannibalism) and reduce their paternity if courtship is brief. We used published data to infer that the female's threshold is approximately 100 min of courtship. We support this hypothesis by showing that premature cannibalism is common when the male's courtship duration is below this threshold, but is infrequent and unrelated to duration once courtship exceeds the threshold. We then ask whether females discriminate the source of courtship when rival males compete, as this will determine the effect of the threshold on male competitive tactics. We staged competitions where 'resident' males initially courted females in the absence of competition, exceeding the courtship threshold before 'intruding' males were introduced. Intruding males mated rapidly but were not prematurely cannibalized by females, in contrast to cases where competition starts before the threshold is surpassed. This suggests females do not distinguish which male satisfies the threshold, allowing intruders to parasitize the courtship efforts of residents. To our knowledge, such exploitation of mating efforts by rival males mediated by a female choice threshold has not been demonstrated elsewhere. Ironically, this female choice threshold and the attendant possibility of courtship parasitism may lead to selection for lower-quality males to recognize and seek out (rather than avoid) webs in which competitors are already present.

A test of ecological and sexual selection hypotheses for geographical variation in coloration and morphology of golden‐crowned kinglets (Regulus satrapa)

AbstractAim To date, studies on geographical variation have extensively investigated Bergmann’s rule, yet Gloger’s rule remains infrequently tested, and climatic predictors of variation in carotenoid coloration have not yet been studied. In addition, hypotheses based on sexual selection, which predict that sexual dimorphism should vary with population density and climatic conditions, have received little attention. Our goals were to characterize geographical variation in the coloration and morphology of golden‐crowned kinglets, Regulus satrapa (Passeriformes, Regulidae), and to investigate possible ecological and sexual selection correlates of this variation.Location The entire species range of golden‐crowned kinglets, comprising North and Central America.Methods We collected data from 511 museum specimens, dating from 1847 to 2006, encompassing all five subspecies of golden‐crowned kinglets. We used reflectance spectrometry to quantify crown and mantle coloration, and measured wing‐chord and tarsus length to approximate body size. We obtained geographical and climatic data from online databases, and population density estimates from the literature.Results There were significant subspecific and gender differences in crown coloration and morphology: male kinglets were generally larger and more colourful. Our data revealed mixed support for Bergmann’s rule: tarsus length decreased with increasing latitude, while patterns of variation in wing‐chord and tarsus length showed conflicting results with temperature. Mantle coloration exhibited an opposite trend to that predicted by Gloger’s rule: upper parts became lighter with increasing relative humidity. Crown coloration was negatively correlated with actual evapotranspiration, suggesting that levels of primary productivity are not directly linked to carotenoid abundance. Sexual dimorphism and dichromatism generally increased with greater population density, lower latitudes and elevations, and warmer temperatures, supporting a previously observed pattern of variation in sexual dimorphism.Main conclusions Geographical variation in golden‐crowned kinglets yielded mixed support for Bergmann’s rule and contradicted Gloger’s rule, suggesting that other mechanisms may be operating. Allen’s rule is likely to be a stronger factor influencing tarsus length. Differences in the degree of sexual dimorphism and dichromatism in varying climatic conditions suggest that the intensity of sexual selection differs between habitats. Further studies on geographical variation in sexual dimorphism in various taxa may reveal a previously unrecognized ecogeographical rule.

Fluctuating mate preferences in a marine fish

According to theory, directional female choice for male sexual ornaments is expected to erode underlying genetic variation. Considerable attention, in this regard, has been given to understanding the ubiquity of heritable genetic variation in both female choice and male sexual traits. One intriguing possibility emerging from this work is that persistent genetic variation could be maintained, over time, by variation in female mate preferences. Here, we report the results of a four-year study showing significant year-to-year fluctuations in mate preferences in a small marine fish, the sand goby, Pomatoschistus minutus. Although the average size of mature fish varied across years, we were unable to find direct evidence linking this variation to differences in female preferences among years. Our results, nevertheless, underscore the importance of temporal fluctuations in female mate preferences, as these can have important consequences for understanding variation in sexual traits and the intensity of sexual selection.

Comparisons on Reproductive and Aggressive Behaviors in Male Tibetan Macaques (<I>Macaca thibetana</I>) of Two Different Operational Sex Ratio Troops at Huangshan, China

Operational sex ratio (OSR) is the ratio of the number of sexually mature males to the estrous females, which is used to predict the intensity of sexual selection. During the mating season (August－December 2007), we studied reproductive and aggressive behaviors in sexually mature male Tibetan macaques (Macaca thibitana) in two troops (YA₁ and YA₂) at Huangshan, China. The behavioral data were collected by focal animal sampling, ad libitum sampling and continuous recording. The purpose is to discuss the impact of OSR on mating competition of male Tibetan macaques. The OSR of YA₁ was 0.4:1, and the OSR of YA₂ was 0.9:1. The male-biased in OSR of YA₂ was higher than YA₁. The sexually mature males of high rank in YA₁ troop showed higher rates of reproductive behaviors than the males in YA₂: genital-inspecting, grimace, sexual chase and mating, respectively (P＜0.01). The sexually mature males of middle rank in YA₁ troop showed higher rates of reproductive behaviors than the males in YA₂: genital-inspecting, grimace, sexual chase (P＜0.01). The sexually mature males of middle rank in YA₁ troop also showed higher rate of mating behavior than the males in YA₂ (P＜0.05). However, there was no significant difference in reproductive behaviors between sexually mature males of low rank in two troops. The sexually mature males of high rank in YA₂ troop showed higher rate of aggressive behavior than the males in YA₁(P＜0.01). The sexually mature males of middle rank in YA₂ troop showed higher rate of aggressive behavior than the males in YA₁(P＜0.05). The sexually mature males of low rank in YA₁ troop didn’t show any aggressive behavior. These results suggested that the intensification of male-biased in OSR can lead to the decrease of reproductive behaviors and the increase of aggressive behavior of male Tibetan macaques. The intensity of mating competition of sexually mature males in YA₂ troop was stronger than the males in YA₁. Our results support the prediction of the impact of OSR on mating competition by sexual selection theory.

Hidden dichromatism in the Burrowing Parrot (Cyanoliseus patagonus) as revealed by spectrometric colour analysis

Bird colour perception differs fundamentally from that of humans. Birds have more cone types in the retina, including UV or violet cones, which enable them to perceive a wider spectral range. Thus, human colour perception can be deceiving when assessing functional aspects of bird plumage coloration, such as the intensity of sexual selection. In this study we measured reflectance spectra of different plumage regions of male and female Burrowing Parrot (Cyanoliseus patagonus) individuals. Although not obvious to human eyes, spectrometry revealed that adults are sexually dichromatic. Plumage regions with structure-based (blue) and structure-psittacofulvin pigment-based (green) coloration differed in achromatic brightness. In contrast, the psittacofulvin- based (red) region differed in spectral shape between the sexes. Thus, Burrowing Parrot is among the growing number of bird species which were formerly classed as sexually monochro- matic based on human vision, but which are actually sexually dichromatic.

Hidden dichromatism in the Burrowing Parrot (Cyanoliseus patagonus) as revealed by spectrometric colour analysis

Bird colour perception differs fundamentally from that of humans. Birds have more cone types in the retina, including UV or violet cones, which enable them to perceive a wider spectral range. Thus, human colour perception can be deceiving when assessing functional aspects of bird plumage coloration, such as the intensity of sexual selection. In this study we measured reflectance spectra of different plumage regions of male and female Burrowing Parrot (Cyanoliseus patagonus) individuals. Although not obvious to human eyes, spectrometry revealed that adults are sexually dichromatic. Plumage regions with structure-based (blue) and structure-psittacofulvin pigment-based (green) coloration differed in achromatic brightness. In contrast, the psittacofulvin-based (red) region differed in spectral shape between the sexes. Thus, Burrowing Parrot is among the growing number of bird species which were formerly classed as sexually monochromatic based on human vision, but which are actually sexually dichromatic.

The allometric pattern of sexually size dimorphic feather ornaments and factors affecting allometry

The static allometry of secondary sexual characters is currently subject to debate. While some studies suggest an almost universal positive allometry for such traits, but isometry or negative allometry for nonornamental traits, other studies maintain that any kind of allometric pattern is possible. Therefore, we investigated the allometry of sexually size dimorphic feather ornaments in 67 species of birds. We also studied the allometry of female feathers homologous to male ornaments (female ornaments in the following) and ordinary nonsexual traits. Allometries were estimated as reduced major axis slopes of trait length on tarsus length. Ornamental feathers showed positive allometric slopes in both sexes, although that was not a peculiarity for ornamental feathers, because nonsexual tail feathers also showed positive allometry. Migration distance (in males) and relative size of the tail ornament (in females) tended to be negatively related to the allometric slope of tail feather ornaments, although these results were not conclusive. Finally, we found an association between mating system and allometry of tail feather ornaments, with species with more intense sexual selection showing a smaller degree of allometry of tail ornaments. This study is consistent with theoretical models that predict no specific kind of allometric pattern for sexual and nonsexual characters.

Mate choice and sexual selection: What have we learned since Darwin?

Charles Darwin laid the foundation for all modern work on sexual selection in his seminal book The Descent of Man, and Selection in Relation to Sex. In this work, Darwin fleshed out the mechanism of sexual selection, a hypothesis that he had proposed in The Origin of Species. He went well beyond a simple description of the phenomenon by providing extensive evidence and considering the far-reaching implications of the idea. Here we consider the contributions of Darwin to sexual selection with a particular eye on how far we have progressed in the last 150 years. We focus on 2 key questions in sexual selection. First, why does mate choice evolve at all? And second, what factors determine the strength of mate choice (or intensity of sexual selection) in each sex? Darwin provided partial answers to these questions, and the progress that has been made on both of these topics since his time should be seen as one of the great triumphs of modern evolutionary biology. However, a review of the literature shows that key aspects of sexual selection are still plagued by confusion and disagreement. Many of these areas are complex and will require new theory and empirical data for complete resolution. Overall, Darwin's contributions are still surprisingly relevant to the modern study of sexual selection, so students of evolutionary biology would be well advised to revisit his works. Although we have made significant progress in some areas of sexual selection research, we still have much to accomplish.

Sexually dimorphic immune response in the harem polygynous Wellington tree weta Hemideina crassidens

Abstract Adult males are often less immunocompetent than females. One explanation for this is that intense sexual selection causes males to trade‐off investment in immunity with traits that increase mating success. This hypothesis is tested in the Wellington tree weta (Hemideina crassidens), a large, sexually dimorphic orthopteran insect in which males possess enormous mandibular weaponry used during fights for access to female mates. Field‐collected males have a significantly greater immune response (greater melanotic encapsulation) than females, suggesting that body condition, longevity or an allied trait is important to male fitness, or that females require materials for egg production that would otherwise be used to boost immunity. Although immunity is expected to trade‐off against reproductive traits in both sexes, there is no significant relationship between immune response and weapon or testes size in males, nor fecundity in females.

Sexual selection drives weak positive selection in protamine genes and high promoter divergence, enhancing sperm competitiveness

Phenotypic adaptations may be the result of changes in gene structure or gene regulation, but little is known about the evolution of gene expression. In addition, it is unclear whether the same selective forces may operate at both levels simultaneously. Reproductive proteins evolve rapidly, but the underlying selective forces promoting such rapid changes are still a matter of debate. In particular, the role of sexual selection in driving positive selection among reproductive proteins remains controversial, whereas its potential influence on changes in promoter regions has not been explored. Protamines are responsible for maintaining DNA in a compacted form in chromosomes in sperm and the available evidence suggests that they evolve rapidly. Because protamines condense DNA within the sperm nucleus, they influence sperm head shape. Here, we examine the influence of sperm competition upon protamine 1 and protamine 2 genes and their promoters, by comparing closely related species of Mus that differ in relative testes size, a reliable indicator of levels of sperm competition. We find evidence of positive selection in the protamine 2 gene in the species with the highest inferred levels of sperm competition. In addition, sperm competition levels across all species are strongly associated with high divergence in protamine 2 promoters that, in turn, are associated with sperm swimming speed. We suggest that changes in protamine 2 promoters are likely to enhance sperm swimming speed by making sperm heads more hydrodynamic. Such phenotypic changes are adaptive because sperm swimming speed may be a major determinant of fertilization success under sperm competition. Thus, when species have diverged recently, few changes in gene-coding sequences are found, while high divergence in promoters seems to be associated with the intensity of sexual selection.

Does reproductive isolation evolve faster in larger populations via sexually antagonistic coevolution?

Sexual conflict over reproductive investment can lead to sexually antagonistic coevolution and reproductive isolation. It has been suggested that, unlike most models of allopatric speciation, the evolution of reproductive isolation through sexually antagonistic coevolution will occur faster in large populations as these harbour greater levels of standing genetic variation, receive larger numbers of mutations and experience more intense sexual selection. We tested this in bruchid beetle populations (Callosobruchus maculatus) by manipulating population size and standing genetic variability in replicated lines derived from founders that had been released from sexual conflict for 90 generations. We found that after 19 generations of reintroduced sexual conflict, none of our treatments had evolved significant overall reproductive isolation among replicate lines. However, as predicted, measures of reproductive isolation tended to be greater among larger populations. We discuss our methodology, arguing that reproductive isolation is best examined by performing a matrix of allopatric and sympatric crosses whereas measurement of divergence requires crosses with a tester line.

Are parental care trade‐offs in shorebirds driven by parental investment or sexual selection?

Sexual selection, mating systems and parental behaviour are closely linked, although the exact nature of their relationship is controversial. The parental investment hypothesis (PIH) states that parental care disparity drives sexual selection intensity, because the sex providing less care competes for the sex that provides more. In contrast, the sexual selection hypothesis (SSH) asserts that more intense sexual selection on males leads to reduced male parental investment. We tested these hypotheses using directional phylogenetic comparative methods in shorebirds, which have an unusually diverse array of breeding systems. Changes in parental care and sexual selection intensity were tightly correlated, and we carried out three sets of analyses focusing on changes in male behaviour, female behaviour and in either sex. The results from the analyses were consistent with both PIH and SSH, although the patterns in male transition were sensitive to model values. We propose two explanations for these results. First, phylogenetic transitions may be idiosyncratic so that they depend on the ecological circumstances of individual species. Second, transitions in social traits, such as breeding systems, may be rapid and take place in ecological time, so directional phylogenetic methods that work through longer time scales may not infer accurately the timing and direction of all changes.

A TEST AND REVIEW OF THE ROLE OF EFFECTIVE POPULATION SIZE ON EXPERIMENTAL SEXUAL SELECTION PATTERNS

Experimental evolution, particularly experimental sexual selection in which sexual selection strength is manipulated by altering the mating system, is an increasingly popular method for testing evolutionary theory. Concerns have arisen regarding genetic diversity variation across experimental treatments: differences in the number and sex ratio of breeders (effective population size; N(e)) and the potential for genetic hitchhiking, both of which may cause different levels of genetic variation between treatments. Such differences may affect the selection response and confound interpretation of results. Here we use both census-based estimators and molecular marker-based estimates to empirically test how experimental evolution of sexual selection in Drosophila pseudoobscura impacts N(e) and autosomal genetic diversity. We also consider effects of treatment on X-linked N(e)s, which have previously been ignored. Molecular autosomal marker-based estimators indicate that neither N(e) nor genetic diversity differs between treatments experiencing different sexual selection intensities; thus observed evolutionary responses reflect selection rather than any confounding effects of experimental design. Given the increasing number of studies on experimental sexual selection, we also review the census N(e)s of other experimental systems, calculate X-linked N(e), and compare how different studies have dealt with the issues of inbreeding, genetic drift, and genetic hitchhiking to help inform future designs.

Secondary Sexual Traits and Individual Phenotype in Male Green Frogs (Rana clamitans)

Secondary sexual characteristics are often under intense sexual selection as ornaments for attracting mates or as traits associated with male-male competition such as combat. Secondary sexual traits under sexual selection are predicted to be related to male phenotype, because only high-quality males should be capable of developing these costly and exaggerated traits. Therefore, high-quality males should have larger or more elaborate secondary sexual traits than poor-quality males. We used the Green Frog (Rana clamitans) to test this hypothesis using three sexually dimorphic traits (the enlarged forelimbs, enlarged tympanum, and yellow throat of the male Green Frog) and two measures of male phenotype (body condition and size). Forelimb size was condition-dependent—males with larger forelimbs were in better condition than males with small forelimbs. Males with an intermediate shade of yellow on their throat tended to be in better condition than males with dark and light yellow throats, but the degree of yellow was related to body size—males with a dark yellow throat were larger than males with a light yellow throat. There was no evidence of condition-dependence of tympanum diameter; although after correcting for body size, males with a large tympanum had large forelimbs suggesting that tympanum size reflected some other, unmeasured, indicator of male quality. These results are consistent with the hypothesis that males produce multiple signals and cues that reflect different aspects of male quality. Females can maximize the benefits of mate choice by using multiple traits to assess male quality.

Why men have shorter lives than women: Effects of resource availability, infectious disease, and senescence

Senescence arises from age-specific deterioration of the soma as a consequence of optimization of life history, and such effects of senescence should appear when comparing species that differ in intensity of sexual selection, as well as when comparing, within a species, the two sexes that often differ in intensity of sexual selection. However, any extrinsic cause of mortality that reduces life expectancy will reduce the possibility of detecting sex-specific differences in senescence. We investigated geographical variation in human sex differences in longevity across 121 countries to test whether differences in sexual competition for limiting resources, reflecting intensity of sexual selection, affected sex differences in longevity. Men on average lived 5 years shorter than women. High rates of childhood morbidity and mortality reduced the sex difference in longevity, while increased overall longevity increased the sex difference in longevity. Increased resource availability estimated from gross domestic product per capita reduced the sex difference in longevity, accounting for 10% of the variance, while there was no additional effect of income inequality as reflected by the Gini coefficient. In a separate analysis of sex differences in longevity among the states of the US, there was a strong effect of the Gini coefficient on sex difference in longevity, with the negative effect on male longevity being stronger than that on female longevity. In contrast, there was only a marginal effect of average household income. Thus, there was evidence of increased competition for resources contributing to increased sex differences in longevity within a single nation.

Sexual selection on cuticular hydrocarbons in the Australian field cricket, Teleogryllus oceanicus

BackgroundFemales in a wide range of taxa have been shown to base their choice of mates on pheromone signals. However, little research has focussed specifically on the form and intensity of selection that mate choice imposes on the pheromone signal. Using multivariate selection analysis, we characterise directly the form and intensity of sexual selection acting on cuticular hydrocarbons, chemical compounds widely used in the selection of mates in insects. Using the Australian field cricket Teleogryllus oceanicus as a model organism, we use three measures of male attractiveness to estimate fitness; mating success, the duration of courtship required to elicit copulation, and subsequent spermatophore attachment duration.ResultsWe found that all three measures of male attractiveness generated sexual selection on male cuticular hydrocarbons, however there were differences in the form and intensity of selection among these three measures. Mating success was the only measure of attractiveness that imposed both univariate linear and quadratic selection on cuticular hydrocarbons. Although we found that all three attractiveness measures generated nonlinear selection, again only mating success was found to exert statistically significant stabilizing selection.ConclusionThis study shows that sexual selection plays an important role in the evolution of male cuticular hydrocarbon signals.

Social fishes and single mothers: brain evolution in African cichlids

As with any organ, differences in brain size--after adequate control of allometry--are assumed to be a response to selection. With over 200 species and an astonishing diversity in niche preferences and social organization, Tanganyikan cichlids present an excellent opportunity to study brain evolution. We used phylogenetic comparative analyses of sexed adults from 39 Tanganyikan cichlid species in a multiple regression framework to investigate the influence of ecology, sexual selection and parental care patterns on whole brain size, as well as to analyse sex-specific effects. First, using species-specific measures, we analysed the influence of diet, habitat, form of care (mouthbrooding or substrate guarding), care type (biparental or female only) and intensity of sexual selection on brain size, while controlling for body size. Then, we repeated the analyses for male and female brain size separately. Type of diet and care type were significantly correlated with whole brain size. Sex-specific analyses showed that female brain size correlated significantly with care type while male brain size was uncorrelated with care type. Our results suggest that more complex social interactions associated with diet select for larger brains and further that the burden of uniparental care exerts high cognitive demands on females.

Sexual Selection for Male Mobility in a Giant Insect with Female‐Biased Size Dimorphism

Female-biased size dimorphism, in which females are larger than males, is prevalent in many animals. Several hypotheses have been developed to explain this pattern of dimorphism. One of these hypotheses, the mobility hypothesis, suggests that female-biased size dimorphism arises because smaller males are favored in scramble competition for mates. Using radiotelemetry, we assessed the mobility hypothesis in the Cook Strait giant weta (Deinacrida rugosa), a species with strong female-biased size dimorphism, and tested the prediction that male traits promoting mobility (i.e., longer legs, smaller bodies) are useful in scramble competition for mates and thus promote reproductive success. Our predictions were supported: males with longer legs and smaller bodies exhibited greater mobility (daily linear displacement when not mating), and more mobile males had greater insemination success. No phenotypic traits predicted female mobility or insemination success. In species with female-biased size dimorphism, sexual selection on males is often considered to be weak compared to species in which males are large or possess weaponry. We found that male giant weta experience sexual selection intensities on par with males of a closely related harem-defending polygynous species, likely because of strong scramble competition with other males.

Ecological gradient of sexual selection: elevation and song elaboration in finches

Ecological gradients in natural and sexual selection often result in evolutionary diversification of morphological, life history, and behavioral traits. In particular, elevational changes in habitat structure and climate not only covary with intensity of sexual selection in many taxa, but may also influence evolution of mating signals. Here we examined variation in courtship song in relation to elevation of breeding across cardueline finches-a subfamily of birds that occupies the widest elevational range of extant birds and shows extensive variation in life histories and sexual selection along this range. We predicted that decrease in sexual selection intensity with elevation of breeding documented in this clade would result in a corresponding evolutionary reduction in elaboration of courtship songs. We controlled for the effects of phylogeny, morphology, and habitat structure to uncover a predicted elevational decline in courtship song elaboration; species breeding at lower elevations sang more elaborated and louder songs compared to their sister species breeding at higher elevations. In addition, lower elevation species had longer songs with more notes, whereas frequency components of song did not vary with elevation. We suggest that changes in sexual selection account for the observed patterns of song variation and discuss how elevational gradient in sexual selection may facilitate divergence in mating signals potentially reinforcing or promoting speciation.

EXPERIMENTAL EVIDENCE THAT SEXUAL CONFLICT INFLUENCES THE OPPORTUNITY, FORM AND INTENSITY OF SEXUAL SELECTION

Sexual interactions are often rife with conflict. Conflict between members of the same sex over opportunities to mate has long been understood to effect evolution via sexual selection. Although conflict between males and females is now understood to be widespread, such conflict is seldom considered in the same light as a general agent of sexual selection. Any interaction between males or females that generates variation in fitness, whether due to conflict, competition or mate choice, can potentially influence sexual selection acting on a range of male traits. Here we seek to address a lack of direct experimental evidence for how sexual conflict influences sexual selection more broadly. We manipulate a major source of sexual conflict in the black field cricket, Teleogryllus commodus, and quantify the resulting changes in the nature of sexual selection using formal selection analysis to statistically compare multivariate fitness surfaces. In T. commodus, sexual conflict occurs over the attachment time of an external spermatophore. By experimentally manipulating the ability of males and females to influence spermatophore attachment, we found that sexual conflict significantly influences the opportunity, form, and intensity of sexual selection on male courtship call and body size. When males were able to harass females, the opportunity for selection was smaller, the form of selection changed, and sexual selection was weaker. We discuss the broader evolutionary implications of these findings, including the contributions of sexual conflict to fluctuating sexual selection and the maintenance of additive genetic variation.